A Meta-Analysis of Thyroid-Related Traits Reveals Novel Loci and Gender-Specific Differences in the Regulation of Thyroid Function

• Eleonora Porcu ,

  Contributed equally to this work with: Eleonora Porcu, Marco Medici, Giorgio Pistis

  Affiliations Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, c/o Cittadella Universitaria di Monserrato, Monserrato, Cagliari, Italy, Dipartimento di Scienze Biomediche, Università di Sassari, Sassari, Italy

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• Marco Medici ,

  Contributed equally to this work with: Eleonora Porcu, Marco Medici, Giorgio Pistis

  Affiliation Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands

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• Giorgio Pistis ,

  Contributed equally to this work with: Eleonora Porcu, Marco Medici, Giorgio Pistis

  Current address: Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, c/o Cittadella Universitaria di Monserrato, Monserrato, Cagliari, Italy; Dipartimento di Scienze Biomediche, Università di Sassari, Sassari, Italy

  Affiliations Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano, Italy, Università degli Studi di Trieste, Trieste, Italy

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• Claudia B. Volpato,

  Affiliation Center for Biomedicine, European Academy Bozen/Bolzano (EURAC), Bolzano, Italy (Affiliated Institute of the University of Lübeck, Lübeck, Germany)

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• Scott G. Wilson,

  Affiliations Department of Endocrinology and Diabetes, Sir Charles Gairdner Hospital, Nedlands, Western Australia, Australia, Department of Twin Research and Genetic Epidemiology, King's College London, London, United Kingdom, School of Medicine and Pharmacology, University of Western Australia, Crawley, Western Australia, Australia

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• Anne R. Cappola,

  Affiliation University of Pennsylvania School of Medicine, Philadelphia, Pennsylvania, United States of America

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• Steffan D. Bos,

  Affiliations Leiden University Medical Center, Molecular Epidemiology, Leiden, The Netherlands, Netherlands Consortium for Healthy Ageing, Leiden, The Netherlands

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• Joris Deelen,

  Affiliations Leiden University Medical Center, Molecular Epidemiology, Leiden, The Netherlands, Netherlands Consortium for Healthy Ageing, Leiden, The Netherlands

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• Martin den Heijer,

  Affiliations Department of Endocrinology, Radboud University Nijmegen Medical Centre, Nijmegen, The Netherlands, Department of Internal Medicine, Free University Medical Center, Amsterdam, The Netherlands

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• Rachel M. Freathy,

  Affiliation Genetics of Complex Traits, Peninsula College of Medicine and Dentistry, University of Exeter, Exeter, United Kingdom

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• Jari Lahti,

  Affiliation Institute of Behavioural Sciences, University of Helsinki, Helsinki, Finland

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• Chunyu Liu,

  Affiliation Center for Population Studies, National Heart, Lung, and Blood Institute, Framingham, Massachusetts, United States of America

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• Lorna M. Lopez,

  Affiliations Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, United Kingdom, Department of Psychology, University of Edinburgh, Edinburgh, United Kingdom

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• Ilja M. Nolte,

  Affiliation Unit of Genetic Epidemiology and Bioinformatics, Department of Epidemiology, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands

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• Jeffrey R. O'Connell,

  Affiliation Department of Medicine, University of Maryland Medical School, Baltimore, Maryland, United States of America

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• Toshiko Tanaka,

  Affiliation Clinical Research Branch, National Institute on Aging, Baltimore, Maryland, United States of America

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• Stella Trompet,

  Affiliations Department of Cardiology, Leiden University Medical Center, Leiden, The Netherlands, Department of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, The Netherlands

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• Alice Arnold,

  Affiliation Cardiovascular Health Research Unit and Department of Medicine, University of Washington, Seattle, Washington, United States of America

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• Stefania Bandinelli,

  Affiliation Geriatric Unit, Azienda Sanitaria Firenze (ASF), Florence, Italy

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• Marian Beekman,

  Affiliations Leiden University Medical Center, Molecular Epidemiology, Leiden, The Netherlands, Netherlands Consortium for Healthy Ageing, Leiden, The Netherlands

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• Stefan Böhringer,

  Affiliation Leiden University Medical Center, Medical Statistics and Bioinformatics, Leiden, The Netherlands

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• Suzanne J. Brown,

  Affiliation Department of Endocrinology and Diabetes, Sir Charles Gairdner Hospital, Nedlands, Western Australia, Australia

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• Brendan M. Buckley,

  Affiliation Department of Pharmacology and Therapeutics, University College Cork, Cork, Ireland

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• Clara Camaschella,

  Affiliations Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano, Italy, Vita e Salute University, San Raffaele Scientific Institute, Milano, Italy

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• Anton J. M. de Craen,

  Affiliation Department of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, The Netherlands

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• Gail Davies,

  Affiliation Department of Psychology, University of Edinburgh, Edinburgh, United Kingdom

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• Marieke C. H. de Visser,

  Affiliation Department for Health Evidence, Radboud University Medical Centre, Nijmegen, The Netherlands

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• Ian Ford,

  Affiliation Robertson Center for Biostatistics, University of Glasgow, Glasgow, United Kingdom

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• Tom Forsen,

  Affiliations Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland, Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland, Helsinki University Central Hospital, Unit of General Practice, Helsinki, Finland, Vaasa Health Care Centre, Diabetes Unit, Vaasa, Finland

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• Timothy M. Frayling,

  Affiliation Genetics of Complex Traits, Peninsula College of Medicine and Dentistry, University of Exeter, Exeter, United Kingdom

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• Laura Fugazzola,

  Affiliation Endocrine Unit, Fondazione Ca' Granda Policlinico and Department of Clinical Sciences and Community Health, University of Milan, Milano, Italy

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• Martin Gögele,

  Affiliation Center for Biomedicine, European Academy Bozen/Bolzano (EURAC), Bolzano, Italy (Affiliated Institute of the University of Lübeck, Lübeck, Germany)

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• Andrew T. Hattersley,

  Affiliation Peninsula NIHR Clinical Research Facility, Peninsula College of Medicine and Dentistry, University of Exeter, Exeter, United Kingdom

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• Ad R. Hermus,

  Affiliation Department of Endocrinology, Radboud University Nijmegen Medical Centre, Nijmegen, The Netherlands

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• Albert Hofman,

  Affiliations Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)–sponsored Netherlands Consortium for Healthy Aging (NCHA), Rotterdam, The Netherlands

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• Jeanine J. Houwing-Duistermaat,

  Affiliation Leiden University Medical Center, Medical Statistics and Bioinformatics, Leiden, The Netherlands

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• Richard A. Jensen,

  Affiliation Cardiovascular Health Research Unit and Department of Medicine, University of Washington, Seattle, Washington, United States of America

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• Eero Kajantie,

  Affiliations Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland, Hospital for Children and Adolescents, Helsinki University Central Hospital and University of Helsinki, Helsinki, Finland

  ⨯
• Margreet Kloppenburg,

  Affiliation Department of Clinical Epidemiology and Rheumatology, Leiden University Medical Center, Leiden, The Netherlands

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• Ee M. Lim,

  Affiliations Department of Endocrinology and Diabetes, Sir Charles Gairdner Hospital, Nedlands, Western Australia, Australia, Pathwest Laboratory Medicine WA, Nedlands, Western Australia, Australia

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• Corrado Masciullo,

  Affiliation Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano, Italy

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• Stefano Mariotti,

  Affiliation Dipartimento di Scienze Mediche, Università di Cagliari, c/o Cittadella Universitaria di Monserrato, Monserrato, Cagliari, Italy

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• Cosetta Minelli,

  Affiliation Center for Biomedicine, European Academy Bozen/Bolzano (EURAC), Bolzano, Italy (Affiliated Institute of the University of Lübeck, Lübeck, Germany)

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• Braxton D. Mitchell,

  Affiliation Department of Medicine, University of Maryland Medical School, Baltimore, Maryland, United States of America

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• Ramaiah Nagaraja,

  Affiliation Laboratory of Genetics, National Institute on Aging, Baltimore, Maryland, United States of America

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• Romana T. Netea-Maier,

  Affiliation Department of Endocrinology, Radboud University Nijmegen Medical Centre, Nijmegen, The Netherlands

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• Aarno Palotie,

  Affiliations Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, United Kingdom, Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland, Department of Medical Genetics, University of Helsinki and University Central Hospital, Helsinki, Finland

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• Luca Persani,

  Affiliations Department of Clinical Sciences and Community Health, University of Milan, Milano, Italy, Division of Endocrinology and Metabolic Diseases, IRCCS Ospedale San Luca, Milan, Italy

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• Maria G. Piras,

  Affiliation Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, c/o Cittadella Universitaria di Monserrato, Monserrato, Cagliari, Italy

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• Bruce M. Psaty,

  Affiliations Cardiovascular Health Research Unit, Departments of Medicine, Epidemiology, and Health Services, University of Washington, Seattle, Washington, United States of America, Group Health Research Institute, Group Health Cooperative, Seattle, Washington, United States of America

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• Katri Räikkönen,

  Affiliation Institute of Behavioural Sciences, University of Helsinki, Helsinki, Finland

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• J. Brent Richards,

  Affiliations Department of Twin Research and Genetic Epidemiology, King's College London, London, United Kingdom, Department of Medicine, Jewish General Hospital, McGill University, Montréal, Québec, Canada, Departments of Human Genetics, Epidemiology, and Biostatistics, Jewish General Hospital, Lady Davis Institute, McGill University, Montréal, Québec

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• Fernando Rivadeneira,

  Affiliations Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)–sponsored Netherlands Consortium for Healthy Aging (NCHA), Rotterdam, The Netherlands

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• Cinzia Sala,

  Affiliation Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano, Italy

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• Mona M. Sabra,

  Affiliation Memorial Sloan Kettering Cancer Center, Medicine-Endocrinology, New York, New York, United States of America

  ⨯
• Naveed Sattar,

  Affiliation BHF Glasgow Cardiovascular Research Centre, Faculty of Medicine, Glasgow, United Kingdom

  ⨯
• Beverley M. Shields,

  Affiliation Peninsula NIHR Clinical Research Facility, Peninsula College of Medicine and Dentistry, University of Exeter, Exeter, United Kingdom

  ⨯
• Nicole Soranzo,

  Affiliation Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, United Kingdom

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• John M. Starr,

  Affiliations Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, United Kingdom, Alzheimer Scotland Dementia Research Centre, University of Edinburgh, Edinburgh, United Kingdom

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• David J. Stott,

  Affiliation Academic Section of Geriatric Medicine, Faculty of Medicine, University of Glasgow, Glasgow, United Kingdom

  ⨯
• Fred C. G. J. Sweep,

  Affiliation Department of Laboratory Medicine, Radboud University Nijmegen Medical Center, Nijmegen, The Netherlands

  ⨯
• Gianluca Usala,

  Affiliation Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, c/o Cittadella Universitaria di Monserrato, Monserrato, Cagliari, Italy

  ⨯
• Melanie M. van der Klauw,

  Affiliations LifeLines Cohort Study, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands, Department of Endocrinology, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands

  ⨯
• Diana van Heemst,

  Affiliation Leiden University Medical Center, Gerontology and Geriatrics, Leiden, The Netherlands

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• Alies van Mullem,

  Affiliation Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands

  ⨯
• Sita H.Vermeulen,

  Affiliation Department for Health Evidence, Radboud University Medical Centre, Nijmegen, The Netherlands

  ⨯
• W. Edward Visser,

  Affiliation Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands

  ⨯
• John P. Walsh,

  Affiliations Department of Endocrinology and Diabetes, Sir Charles Gairdner Hospital, Nedlands, Western Australia, Australia, School of Medicine and Pharmacology, University of Western Australia, Crawley, Western Australia, Australia

  ⨯
• Rudi G. J. Westendorp,

  Affiliation Leiden University Medical Center, Gerontology and Geriatrics, Leiden, The Netherlands

  ⨯
• Elisabeth Widen,

  Affiliation Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland

  ⨯
• Guangju Zhai,

  Affiliations Department of Twin Research and Genetic Epidemiology, King's College London, London, United Kingdom, Discipline of Genetics, Faculty of Medicine, Memorial University of Newfoundland, St. Johns, Newfoundland, Canada

  ⨯
• Francesco Cucca,

  Affiliations Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, c/o Cittadella Universitaria di Monserrato, Monserrato, Cagliari, Italy, Dipartimento di Scienze Biomediche, Università di Sassari, Sassari, Italy

  ⨯
• Ian J. Deary,

  Affiliations Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, United Kingdom, Department of Psychology, University of Edinburgh, Edinburgh, United Kingdom

  ⨯
• Johan G. Eriksson,

  Affiliations Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland, Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland, Helsinki University Central Hospital, Unit of General Practice, Helsinki, Finland, Folkhalsan Research Centre, Helsinki, Finland, Vasa Central Hospital, Vasa, Finland

  ⨯
• Luigi Ferrucci,

  Affiliation Clinical Research Branch, National Institute on Aging, Baltimore, Maryland, United States of America

  ⨯
• Caroline S. Fox,

  Affiliations Division of Intramural Research, National Heart, Lung, and Blood Institute, Framingham, Massachusetts, United States of America, Division of Endocrinology, Hypertension, and Metabolism, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts, United States of America

  ⨯
• J. Wouter Jukema,

  Affiliations Department of Cardiology, Leiden University Medical Center, Leiden, The Netherlands, Durrer Center for Cardiogenetic Research, Amsterdam, The Netherlands, Interuniversity Cardiology Institute of the Netherlands, Utrecht, The Netherlands

  ⨯
• Lambertus A. Kiemeney,

  Affiliations Department for Health Evidence, Radboud University Medical Centre, Nijmegen, The Netherlands, Department of Urology, Radboud University Medical Centre, Nijmegen, The Netherlands

  ⨯
• Peter P. Pramstaller,

  Affiliations Center for Biomedicine, European Academy Bozen/Bolzano (EURAC), Bolzano, Italy (Affiliated Institute of the University of Lübeck, Lübeck, Germany), Department of Neurology, General Central Hospital, Bolzano, Italy, Department of Neurology, University of Lübeck, Lübeck, Germany

  ⨯
• David Schlessinger,

  Affiliation Laboratory of Genetics, National Institute on Aging, Baltimore, Maryland, United States of America

  ⨯
• Alan R. Shuldiner,

  Affiliations Department of Medicine, University of Maryland Medical School, Baltimore, Maryland, United States of America, Geriatric Research and Education Clinical Center, Veterans Administration Medical Center, Baltimore, Maryland, United States of America

  ⨯
• Eline P. Slagboom,

  Affiliations Leiden University Medical Center, Molecular Epidemiology, Leiden, The Netherlands, Netherlands Consortium for Healthy Ageing, Leiden, The Netherlands

  ⨯
• André G. Uitterlinden,

  Affiliations Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands, Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands, Netherlands Genomics Initiative (NGI)–sponsored Netherlands Consortium for Healthy Aging (NCHA), Rotterdam, The Netherlands

  ⨯
• Bijay Vaidya,

  Affiliation Diabetes, Endocrinology and Vascular Health Centre, Royal Devon and Exeter NHS Foundation Trust, Exeter, United Kingdom

  ⨯
• Theo J. Visser,

  Affiliation Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands

  ⨯
• Bruce H. R. Wolffenbuttel,

  Affiliations LifeLines Cohort Study, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands, Department of Endocrinology, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands

  ⨯
• Ingrid Meulenbelt,

  Affiliations Leiden University Medical Center, Molecular Epidemiology, Leiden, The Netherlands, Netherlands Consortium for Healthy Ageing, Leiden, The Netherlands

  ⨯
• Jerome I. Rotter,

  Affiliation Medical Genetics Institute, Cedars-Sinai Medical Center, Los Angeles, California, United States of America

  ⨯
• Tim D. Spector,

  Affiliation Department of Twin Research and Genetic Epidemiology, King's College London, London, United Kingdom

  ⨯
• Andrew A. Hicks,

  Affiliation Center for Biomedicine, European Academy Bozen/Bolzano (EURAC), Bolzano, Italy (Affiliated Institute of the University of Lübeck, Lübeck, Germany)

  ⨯
• Daniela Toniolo,

  Affiliations Division of Genetics and Cell Biology, San Raffaele Research Institute, Milano, Italy, Institute of Molecular Genetics–CNR, Pavia, Italy

  ⨯
• Serena Sanna ,

  * E-mail: serena.sanna@irgb.cnr.it (S Sanna); r.peeters@erasmusmc.nl (RP Peeters); silvia.naitza@irgb.cnr.it (S Naitza)

  ¶These authors also contributed equally to this work.

  Affiliation Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, c/o Cittadella Universitaria di Monserrato, Monserrato, Cagliari, Italy

  ⨯
• Robin P. Peeters ,

  * E-mail: serena.sanna@irgb.cnr.it (S Sanna); r.peeters@erasmusmc.nl (RP Peeters); silvia.naitza@irgb.cnr.it (S Naitza)

  ¶These authors also contributed equally to this work.

  Affiliation Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands

  ⨯
•  [ ... ],
• Silvia Naitza

  * E-mail: serena.sanna@irgb.cnr.it (S Sanna); r.peeters@erasmusmc.nl (RP Peeters); silvia.naitza@irgb.cnr.it (S Naitza)

  ¶These authors also contributed equally to this work.

  Affiliation Istituto di Ricerca Genetica e Biomedica (IRGB), Consiglio Nazionale delle Ricerche, c/o Cittadella Universitaria di Monserrato, Monserrato, Cagliari, Italy

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